A new alvarezsaurid dinosaur (Theropoda, Alvarezsauria) from the Upper Cretaceous Baruungoyot Formation of Mongolia provides insights for bird-like sleeping behavior in non-avian dinosaurs.

Alvarezsauria is a group of early-branching maniraptoran theropods that are distributed globally from the Late Jurassic to the latest Cretaceous. Despite recent increases in the fossil record of this group, the scarcity of complete specimens still restricts interpreting their detailed anatomy, ecology, and evolution. Here, we report a new taxon of derived alvarezsaur, Jaculinykus yaruui gen. et sp. nov., from the Late Cretaceous of Mongolia, which represents a nearly complete and articulated skeleton. Our phylogenetic analysis reveals that Jaculinykus belongs to the sub-clade of Alvarezsauridae, Parvicursorinae, and forms a mononphyletic group with Mononykus and Shuvuuia. Its well-preserved manus has only two fingers, composed of a hypertrophied digit I and greatly reduced digit II, which implies an intermediate condition between the tridactyl manus of Shuvuuia and monodactyl manus of Linhenykus. This highlights a previously unrecognized variation in specialization of alvarezsaurid manus. Notably, the preserved posture of the specimen exhibits a stereotypical avian-like sleeping position seen in the troodontids Mei and Sinornithoides. Evidence of this behavior in the alvarezsaur Jaculinykus suggests that stereotypically avian sleeping postures are a maniraptoran synapomorphy, providing more evidence of bird-like traits being distributed broadly among avian ancestors.

Evolutionary process toward avian-like cephalic thermoregulation system in Theropoda elucidated based on nasal structures.

It has long been discussed whether non-avian dinosaurs were physiologically closer to ectotherms or endotherms, with the internal nasal structure called the respiratory turbinate present in extant endotherms having been regarded as an important clue for this conundrum. However, the physiological function and relevance of this structure for dinosaur physiology are still controversial. Here, we found that the size of the nasal cavity relative to the head size of extant endotherms is larger than those of extant ectotherms, with that of the dromaeosaurid Velociraptor being below the extant endotherms level. The result suggests that a large nasal cavity accommodating a well-developed respiratory turbinate is primarily important as a thermoregulation apparatus for large brains characteristic of endothermic birds and mammals, and the nasal cavity of Velociraptor was apparently not large enough to carry out this role required for an endothermic-sized brain. In addition, a hypothesis that the enlargement of the nasal cavity for brain cooling has been associated with the skull modification in the theropod lineage toward modern birds is proposed herein. In particular, the reduction of the maxilla in derived avialans may have coincided with acquisition of the avian-like cephalic thermoregulation system.

A non-avian dinosaur with a streamlined body exhibits potential adaptations for swimming.

Streamlining a body is a major adaptation for aquatic animals to move efficiently in the water. Whereas diving birds are well known to have streamlined bodies, such body shapes have not been documented in non-avian dinosaurs. It is primarily because most known non-avian theropods are terrestrial, barring a few exceptions. However, clear evidence of streamlined bodies is absent even in the purported semiaquatic groups. Here we report a new theropod, Natovenator polydontus gen. et sp. nov., from the Upper Cretaceous of Mongolia. The new specimen includes a well-preserved skeleton with several articulated dorsal ribs that are posterolaterally oriented to streamline the body as in diving birds. Additionally, the widely arched proximal rib shafts reflect a dorsoventrally compressed ribcage like aquatic reptiles. Its body shape suggests that Natovenator was a potentially capable swimming predator, and the streamlined body evolved independently in separate lineages of theropod dinosaurs.

Unusual pectoral apparatus in a predatory dinosaur resolves avian wishbone homology.

The furcula is a distinctive element of the pectoral skeleton in birds, which strengthens the shoulder region to withstand the rigor of flight. Although its origin among theropod dinosaurs is now well-supported, the homology of the furcula relative to the elements of the tetrapod pectoral girdle (i.e., interclavicle vs clavicles) remains controversial. Here, we report the identification of the furcula in the birdlike theropod Halszkaraptor escuilliei. The bone is unique among furculae in non-avian dinosaurs in bearing a visceral articular facet in the hypocleideal end firmly joined to and overlapped by the sternal plates, a topographical pattern that supports the primary homology of the furcula with the interclavicle. The transformation of the interclavicle into the furcula in early theropods is correlated to the loss of the clavicles, and reinforced the interconnection between the contralateral scapulocoracoids, while relaxing the bridge between the scapulocoracoids with the sternum. The function of the forelimbs in theropod ancestors shifted from being a component of the locomotory quadrupedal module to an independent module specialized to grasping. The later evolution of novel locomotory modules among maniraptoran theropods, involving the forelimbs, drove the re-acquisition of a tighter connection between the scapulocoracoids and the interclavicle with the sternal complex.

A new two-fingered dinosaur sheds light on the radiation of Oviraptorosauria.

Late Cretaceous trends in Asian dinosaur diversity are poorly understood, but recent discoveries have documented a radiation of oviraptorosaur theropods in China and Mongolia. However, little work has addressed the factors that facilitated this diversification. A new oviraptorid from the Late Cretaceous of Mongolia sheds light on the evolution of the forelimb, which appears to have played a role in the radiation of oviraptorosaurs. Surprisingly, the reduced arm has only two functional digits, highlighting a previously unrecognized occurrence of digit loss in theropods. Phylogenetic analysis shows that the onset of this reduction coincides with the radiation of heyuannine oviraptorids, following dispersal from southern China into the Gobi region. This suggests expansion into a new niche in the Gobi region, which relied less on the elongate, grasping forelimbs inherited by oviraptorosaurs. Variation in forelimb length and manus morphology provides another example of niche partitioning in oviraptorosaurs, which may have made possible their incredible diversity in the latest Cretaceous of Asia.

Author Correction: First Ornithomimid (Theropoda, Ornithomimosauria) from the Upper Cretaceous Djadokhta Formation of Tögrögiin Shiree, Mongolia.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

A new alvarezsaurid dinosaur from the Nemegt Formation of Mongolia.

Alvarezsaurid diversity has been markedly increased by recent discoveries from China. However, the number of alvarezsaurid specimens in the Nemegt Formation of Mongolia remained low since the initial report on Mononykus olecranus in 1993. Here we report three new alvarezsaurid specimens from this formation, which were associated with each other and also with multiple oviraptorid skeletons in a small multi-species assemblage. Two of the alvarezsaurid specimens represent a new taxon, Nemegtonykus citus gen. et sp. nov., which is mainly distinguished from other alvarezsaurids by the first sacral vertebra with a subtrapezoidal lamina, the second sacral centrum which is directly co-ossified with ilium, the posterodorsally oriented postacetabular process of ilium, and partial co-ossification between metatarsals II and IV. The other specimen is very similar to M. olecranus in morphology and referred to cf. Mononykus sp. Our phylogenetic analysis recovered Nemegtonykus as a parvicursorine forming a polytomy with several other taxa from the Gobi Desert. The presence of three alvarezsaurid individuals in the same locality indicates that the abundance of alvarezsaurids have been greatly underestimated in the Nemegt dinosaur faunas.

A new hadrosauroid (Dinosauria: Ornithopoda) from the Late Cretaceous Baynshire Formation of the Gobi Desert (Mongolia).

A new genus and species of non-hadrosaurid hadrosauroid, Gobihadros mongoliensis, is described from a virtually complete and undeformed skull and postcranial skeleton, as well as extensive referred material, collected from the Baynshire Formation (Cenomanian-Santonian) of the central and eastern Gobi Desert, Mongolia. Gobihadros mongoliensis is the first non-hadrosaurid hadrosauroid from the Late Cretaceous of central Asia known from a complete, articulated skull and skeleton. The material reveals the skeletal anatomy of a proximate sister taxon to Hadrosauridae in remarkable detail. Gobihadros is similar to Bactrosaurus johnsoni and Gilmoreosaurus mongoliensis, but can be distinguished from them in several autapomorphic traits, including the maximum number (three) of functional dentary teeth per tooth position, a premaxillary oral margin with a 'double-layer morphology', and a sigmoidal dorsal outline of the ilium with a well-developed, fan-shaped posterior process. All of these characters in Gobihadros are inferred to be convergent in Hadrosauridae. Phylogenetic analysis positions Gobihadros mongoliensis as a Bactrosaurus-grade hadrosauromorph hadrosauroid. Its relationship with Maastrichtian hadrosaurids from Asia (e.g., Saurolophus angustirostris, Kerberosaurus manakini, Wulagasaurus dongi, Kundurosaurus nagornyi) are sufficiently distant to indicate that these latter taxa owe their distribution to migration from North America across Beringia, rather than having a common Asian origin with Go. mongoliensis.

A new baby oviraptorid dinosaur (Dinosauria: Theropoda) from the Upper Cretaceous Nemegt Formation of Mongolia.

Recent discoveries of new oviraptorosaurs revealed their high diversity from the Cretaceous Period in Asia and North America. Particularly, at the family level, oviraptorids are among the most diverse theropod dinosaurs in the Late Cretaceous of Mongolia and China. A new oviraptorid dinosaur Gobiraptor minutus gen. et sp. nov. from the Upper Cretaceous Nemegt Formation is described here based on a single holotype specimen that includes incomplete cranial and postcranial elements. The most prominent characters of Gobiraptor are its thickened rostrodorsal end of the mandibular symphysis and a rudimentary lingual shelf on each side of the dentary. Each lingual shelf is lined with small occlusal foramina and demarcated by a weakly developed lingual ridge. This mandibular morphology of Gobiraptor is unique among oviraptorids and likely to be linked to a specialized diet that probably included hard materials, such as seeds or bivalves. The osteohistology of the femur of the holotype specimen indicates that the individual was fairly young at the time of its death. Phylogenetic analysis recovers Gobiraptor as a derived oviraptorid close to three taxa from the Ganzhou region in southern China, but rather distantly related to other Nemegt oviraptorids which, as the results of recent studies, are also not closely related to each other. Gobiraptor increases diversity of oviraptorids in the Nemegt Formation and its presence confirms the successful adaptation of oviraptorids to a mesic environment.

A new iguanodontian (Dinosauria: Ornithopoda) from the Early Cretaceous of Mongolia.

We describe a new iguanodontian ornithopod, Choyrodon barsboldi gen. et sp. nov. from the Albian-aged Khuren Dukh Formation of Mongolia based on several partial skeletons interpreted to represent a subadult growth stage based on osteohistological features. This new taxon is diagnosed by many autapomorphies of the maxilla, nasal, lacrimal, opisthotic, predentary, and surangular. Choyrodon displays an unusual combination of traits, possessing an open antorbital fenestra (a primitive ornithopod trait) together with derived features such as a downturned dentary and enlarged narial fenestra. Histological imaging suggests that the type specimen of Choyrodon would have been a subadult at the time of death. Phylogenetic analysis of two different character matrices do not posit Choyrodon to be the sister taxon or to be more primitive than the iguanodontian Altirhinus kurzanovi, which is found in the same formation. The only resolved relationship of this new taxon is that it was hypothesized to be a sister-taxon with the North American species Eolambia caroljonesa. Though discovered in the same formation and Choyrodon being smaller-bodied than Altirhinus, it does not appear that the former species is an ontogimorph of the latter. Differences in morphology and results of the phylogenetic analyses support their distinction although more specimens of both species will allow better refinement of their uniqueness.

Author Correction: First Ornithomimid (Theropoda, Ornithomimosauria) from the Upper Cretaceous Djadokhta Formation of Tögrögiin Shiree, Mongolia.

A correction to this article has been published and is linked from the HTML and PDF versions of this paper. The error has been fixed in the paper.

Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs.

Maniraptora includes birds and their closest relatives among theropod dinosaurs. During the Cretaceous period, several maniraptoran lineages diverged from the ancestral coelurosaurian bauplan and evolved novel ecomorphologies, including active flight, gigantism, cursoriality and herbivory. Propagation X-ray phase-contrast synchrotron microtomography of a well-preserved maniraptoran from Mongolia, still partially embedded in the rock matrix, revealed a mosaic of features, most of them absent among non-avian maniraptorans but shared by reptilian and avian groups with aquatic or semiaquatic ecologies. This new theropod, Halszkaraptor escuilliei gen. et sp. nov., is related to other enigmatic Late Cretaceous maniraptorans from Mongolia in a novel clade at the root of Dromaeosauridae. This lineage adds an amphibious ecomorphology to those evolved by maniraptorans: it acquired a predatory mode that relied mainly on neck hyperelongation for food procurement, it coupled the obligatory bipedalism of theropods with forelimb proportions that may support a swimming function, and it developed postural adaptations convergent with short-tailed birds.

First Ornithomimid (Theropoda, Ornithomimosauria) from the Upper Cretaceous Djadokhta Formation of Tögrögiin Shiree, Mongolia.

The Upper Cretaceous Djadokhta Formation has been intensively surveyed for its fossil vertebrate fauna for nearly a century. Amongst other theropods, dromaeosaurids and parvicursorines are common in the formation, but ornithomimosaurs are extremely rare. A new ornithomimosaur material was discovered from the Djadokhta Formation, represented by eolian deposits, of the Tögrögiin Shiree locality, Mongolia. This is only the third ornithomimosaur specimen reported from this formation, and includes the astragalus, the calcaneum, the third distal tarsal, and a complete pes. The new material is clearly belonged to Ornithomimidae by its arctometatarsalian foot condition and has the following unique characters; unevenly developed pair of concavities of the third distal tarsal, curved contacts between the proximal ends of second and fourth metatarsals, the elongate fourth digit, and a laterally inclined medial condyle on phalanx IV-1. These diagnostic characters of the Djadokhtan ornithomimosaur indicate that this is a new taxon. Our phylogenetic analysis supports three clades within derived ornithomimosaurs, and the new taxon is placed a member of the derived ornithomimosaurs. The present specimen is the first ornithomimid record from eolian Tögrögiin Shiree locality, and is indicative of their capability to adapt to arid environments.

Oligocene stratigraphy across the Eocene and Miocene boundaries in the Valley of Lakes (Mongolia).

Cenozoic sediments of the Taatsiin Gol and TaatsiinTsagaan Nuur area are rich in fossils that provide unique evidence of mammal evolution in Mongolia. The strata are intercalated with basalt flows. 40Ar/39Ar data of the basalts frame the time of sediment deposition and mammal evolution and enable a composite age chronology for the studied area. We investigated 20 geological sections and 6 fossil localities of Oligocene and early Miocene deposits from this region. Seventy fossil beds yielded more than 19,000 mammal fossils. This huge collection encompasses 175 mammal species: 50% Rodentia, 13% Eulipotyphla and Didelphomorphia, and 12% Lagomorpha. The remaining 25% of species are distributed among herbivorous and carnivorous large mammals. The representation of lower vertebrates and gastropods is comparatively poor. Several hundred SEM images illustrate the diversity of Marsupialia, Eulipotyphla, and Rodentia dentition and give insight into small mammal evolution in Mongolia during the Oligocene and early Miocene. This dataset, the radiometric ages of basalt I (∼31.5 Ma) and basalt II (∼27 Ma), and the magnetostratigraphic data provide ages of mammal assemblages and time ranges of the Mongolian biozones: letter zone A ranges from ∼33 to ∼31.5 Ma, letter zone B from ∼31.5 to ∼28 Ma, letter zone C from ∼28 to 25.6 Ma, letter zone C1 from 25.6 to 24 Ma, letter zone C1-D from 24 to ∼23 Ma, and letter zone D from ∼23 to ∼21 Ma.

Specializations of the mandibular anatomy and dentition of Segnosaurus galbinensis (Theropoda: Therizinosauria).

Definitive therizinosaurid cranial materials are exceptionally rare, represented solely by an isolated braincase and tooth in the North American taxon Nothronychus mckinleyi, the remarkably complete skull of the Asian taxon Erlikosaurus andrewsi, and the lower hemimandibles of Segnosaurus galbinensis. To date, comprehensive descriptions of the former taxa are published; however, the mandibular materials of S. galbinensis have remained largely understudied since their initial description in 1979. Here we provide a comprehensive description of the well-preserved hemimandibles and dentition of S. galbinensis (MPC-D 100/80), from the Upper Cretaceous Bayanshiree Formation, Gobi Desert, Mongolia. The subrectangular and ventrally displaced caudal hemimandible, extreme ventral deflection of the rostral dentary, and edentulism of the caudal dentary of S. galbinensis are currently apomorphic among therizinosaurians. Unique, unreported dental traits including lingually folded mesial carinae, development of a denticulated triangular facet on the distal carinae near the cervix, and extracarinal accessory denticles, suggest a highly specialized feeding strategy in S. galbinensis. The presence of triple carinae on the distalmost lateral tooth crowns is also unique, although may represent an abnormality. Contrasted with the simplistic dentition of the contemporaneous therizinosaurid E. andrewsi, the dentition of S. galbinensis is indicative of niche partitioning in food acquisition, processing, or resources among known therizinosaurids inhabiting Asian ecosystems in the Late Cretaceous. Although not quantitatively correlated with diet, this suite of specializations is otherwise unique among theropod dinosaurs and supports derived inferences of facultative or obligate herbivory in therizinosaurids, ultimately adding novel information to our understanding of ecomorphology in theropods.

Perinatal Specimens of Saurolophus angustirostris (Dinosauria: Hadrosauridae), from the Upper Cretaceous of Mongolia.

BACKGROUND: The Late Cretaceous Nemegt Formation, Gobi Desert, Mongolia has already yielded abundant and complete skeletons of the hadrosaur Saurolophus angustirostris, from half-grown to adult individuals. METHODOLOGY/PRINCIPAL FINDINGS: Herein we describe perinatal specimens of Saurolophus angustirostris, associated with fragmentary eggshell fragments. The skull length of these babies is around 5% that of the largest known S. angustirostris specimens, so these specimens document the earliest development stages of this giant hadrosaur and bridge a large hiatus in our knowledge of the ontogeny of S. angustirostris. CONCLUSIONS/SIGNIFICANCE: The studied specimens are likely part of a nest originally located on a riverbank point bar. The perinatal specimens were buried by sediment carried by the river current presumably during the wet summer season. Perinatal bones already displayed diagnostic characters for Saurolophus angustirostris, including premaxillae with a strongly reflected oral margin and upturned premaxillary body in lateral aspect. The absence of a supracranial crest and unfused halves of the cervical neural arches characterize the earliest stages in the ontogeny of S. angustirostris. The eggshell fragments associated with the perinatal individuals can be referred to the Spheroolithus oogenus and closely resemble those found in older formations (e.g. Barun Goyot Fm in Mongolia) or associated with more basal hadrosauroids (Bactrosaurus-Gilmoreosaurus in the Iren Dabasu Fm, Inner Mongolia, China). This observation suggests that the egg microstructure was similar in basal hadrosauroids and more advanced saurolophines. COMPETING INTERESTS: One of the authors (FE) is employed by the commercial organization Eldonia. Eldonia provided support in the form of a salary for FE, but did not have any additional role or influence in the study design, data collection and analysis, decision to publish, or preparation of the manuscript and it does not alter the authors' adherence to all the PLoS ONE policies on sharing data and materials.

A new mass mortality of juvenile Protoceratops and size-segregated aggregation behaviour in juvenile non-avian dinosaurs.

BACKGROUND: Monodominant bonebeds are a relatively common occurrence for non-avian dinosaurs, and have been used to infer associative, and potentially genuinely social, behavior. Previously known assemblages are characterized as either mixed size-classes (juvenile and adult-sized specimens together) or single size-classes of individuals (only juveniles or only adult-sized individuals within the assemblage). In the latter case, it is generally unknown if these kinds of size-segregated aggregations characterize only a particular size stage or represent aggregations that happened at all size stages. Ceratopsians ("horned dinosaurs") are known from both types of assemblages. METHODS/PRINCIPAL FINDINGS: Here we describe a new specimen of the ceratopsian dinosaur Protoceratops andrewsi, Granger and Gregory 1923 from Mongolia representing an aggregation of four mid-sized juvenile animals. In conjunction with existing specimens of groups of P. andrewsi that includes size-clustered aggregations of young juveniles and adult-sized specimens, this new material provides evidence for some degree of size-clustered aggregation behaviour in Protoceratops throughout ontogeny. This continuity of size-segregated (and presumably age-clustered) aggregation is previously undocumented in non-avian dinosaurs. CONCLUSIONS: The juvenile group fills a key gap in the available information on aggregations in younger ceratopsians. Although we support the general hypothesis that many non-avian dinosaurs were gregarious and even social animals, we caution that evidence for sociality has been overstated and advocate a more conservative interpretation of some data of 'sociality' in dinosaurs.

An exquisitely preserved troodontid theropod with new information on the palatal structure from the Upper Cretaceous of Mongolia.

Troodontidae is a clade of small-bodied theropod dinosaurs. A new troodontid, Gobivenator mongoliensis gen. et sp. nov., is described based on the most complete skeleton of a Late Cretaceous member of this clade presently known, from the Campanian Djadokhta Formation in the central Gobi Desert. G. mongoliensis is different from other troodontids in possessing a pointed anterior end of the fused parietal and a fossa on the surangular in front of the posterior surangular foramen. The skull was superbly preserved in the specimen and provides detailed information of the entire configuration of the palate in Troodontidae. Overall morphology of the palate in Gobivenator resembles those of dromaeosaurids and Archaeopteryx, showing an apparent trend of elongation of the pterygoid process of the palatine and reduction of the pterygopalatine suture toward the basal Avialae. The palatal configuration suggests that the skull of Gobivenator would have been akinetic but had already acquired prerequisites for later evolution of cranial kinesis in birds, such as the loss of the epipterygoid and reduction in contact areas among bones.